Sperm Competition and Its Evolutionary Consequences in the Insects

Leigh W. Simmons is an Australian Research Council Senior Research Fellow and Research Professor in the Department of Zoology. University of Western Australia. He is an Associate Editor for Bahavioral Ecology and Sociobiology and Executive Editor of Animal Behavior.

• Preface ix Dedication and Acknowledgments xv Chapter 1: Sexual Selection and Sperm Competition 1 1.1 Sexual Selection 1 1.2 Sexual Differences and the Evolution of Anisogamy 5 1.3 Sperm Competition and Sexual Selection 10 1.4 Summary 20 Chapter 2: Sperm Utilization: Concepts, Patterns, and Processes 22 2.1 Introduction 22 2.2 Classification and Definition of Terms Used in the Sperm Competition Literature 23 2.3 Quantifying Paternity 29 2.4 Patterns of Sperm Utilization: An Overview 37 2.5 Mechanisms 40 2.6 Sperm Utilization and Multiple Mating 53 2.7 Sperm Utilization in Natural Populations 55 2.8 Mechanisms and the Potential for Selection 56 2.9 Summary 60 Chapter 3: Avoidance of Sperm Competition I: Morphological Adaptations 69 3.1 Introduction 69 3.2 Internal Fertilization 69 3.3 Sperm Removal and Repositioning 70 3.4 Alternative Explanations for Complex Genitalia 82 3.5 Summary 89 Chapter 4: Avoidance of Sperm Competition II: Physiological Adaptations 91 4.1 Introduction 91 4.2 Mating Plugs 97 4.3 Seminal Products 114 4.4 Pheromones 141 4.5 Summary 142 4.6 Avoidance of Sperm Chapter 5: Competition III: Behavioral Adaptations 144 5.1 Introduction 144 5.2 Theoretical Models of Mate Guarding 146 5.3 Evidence for Mate Guarding in Insects 149 5.4 Alternative Hypotheses 168 5.5 Adaptations for Efficient Guarding 179 5.6 Male Mate Choice 183 5.7 Summary 186 Chapter 6: Copula Duration 198 6.1 Introduction 198 6.2 Sperm Displacement: Optimal Copula Duration in Dung Flies 202 6.3 Copula Duration with Sperm Mixing 212 6.4 Evidence Consistent with an Optimization of Copula Duration 213 6.5 Female Influences 218 6.6 Alternative Explanations 220 6.7 Summary 221 Chapter 7: Sperm in Competition I: Strategic Ejaculation 223 7.1 Introduction 223 7.2 Sperm Competition Games 227 7.3 Cryptic Male Choice 245 7.4 Cryptic Female Choice 247 7.5 Summary 249 Chapter 8: Sperm in Competition II: Sperm Morphology 250 8.1 Introduction 250 8.2 Sperm Size 252 8.3 Sperm Polymorphism 263 8.4 Intraejaculate Sperm Competition 274 8.5 Summary 275 Chapter 9: Ejaculate Manipulation: Mechanisms of Female Choice 277 9.1 Introduction 277 9.2 Influence over Remating 279 9.3 Influence over Sperm Transfer 282 9.4 Influence over Sperm Storage 288 9.5 Sperm Selection 299 9.6 Summary 317 Chapter 10: Social insects 319 10.1 Introduction 319 10.2 Multiple Mating and Multiple Paternity 320 10.3 Social Consequences of Sperm Competition 325 10.4 Alternative Hypotheses for Multiple Mating 334 10.5 Summary 341 Chapter 11: Broader Significance 349 11.1 Sperm Competition and Sexual Selection 349 11.2 Life History Evolution 350 11.3 Speciation 354 11.4 Concluding Remarks 356 References 357 Taxonomic Index 427 Subject Index 432

"Leigh Simmons is better qualified than most to define the current state of sperm competition. This book is an exellent summary of a recent, important, and relatively large addition to our understanding of evolution and its consequences. It is an essential read for anyone interested in the reproductive determinates of fitness."--Michael T. Siva-Jothy, Nature "An introductory chapter clearly presents fundamental concepts and terms and general methods used for quantifying sperm competition... Numerous figures and tables enhance the value of this work, which includes more than 1,000 references. Highly recommended for evolutionary biologists, behavioral ecologists, and entomologists."--Choice "This well-illustrated, heavily referenced book will be valuable to researches interested in the evolution of reproductive biology, no matter what group they study."--John Alcock, The Quarterly Review of Biology